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Do mass extinctions grade continuously into the background extinctions occurring throughout the history of life, or are they a fundamentally distinct phenomenon that cannot be explained by processes responsible for background extinction? Various criteria have been proposed for addressing this question, including approaches based on physical mechanisms, ecological selectivity, and statistical characterizations of extinction intensities.
Here I propose a framework defining three types of continuity of mass and background extinctions—continuity of cause, continuity of effect, and continuity of magnitude. I test the third type of continuity with a statistical method based on kernel density estimation. Previous statistical approaches typically have examined quantitative characteristics of mass extinctions (such as metrics of extinction intensity) and compared them with the distribution of such characteristics associated with background extinctions. If mass extinctions are outliers, or are separated by a gap from background extinctions, the distinctness of mass extinctions is supported.
In this paper I apply Silverman's Critical Bandwidth Test to test for the continuity of mass extinctions by applying kernel density estimation and bootstrap modality testing. The method improves on existing work based on searching for gaps in histograms, in that it does not depend on arbitrary choices of parameters (such as bin widths for histograms), and provides a direct estimate of the significance of continuities or gaps in observed extinction intensities. I am thus able to test rigorously whether differences between mass extinctions and background extinctions are statistically significant.
I apply the methodology to Sepkoski's database of Phanerozoic marine genera. I conclude that mass and background extinctions appear to be continuous at this third level—continuity of magnitude—even though evidence suggests that they are discontinuous at the first and second levels.
A drawback to most existing methods of calculating confidence limits on fossil ranges is their assumption that the probability of collecting a taxon through a stratigraphic section is constant. Marshall (1997) described an approach that would circumvent this problem, but it requires knowing the probability of collection as a function of stratigraphic position. Multivariate paleoecological methods, such as detrended correspondence analysis (DCA), offer a means of estimating these probabilities. DCA axis 1 sample scores can be used to quantify facies change through a stratigraphic section, and to calculate the probability of collection of a taxon relative to DCA axis 1. From these two, the probability of collection of each taxon can be estimated for each horizon in the measured section. This approach is applied here to the Upper Ordovician Kope Formation of the Cincinnati, Ohio, area to distinguish between disappearances of taxa that are driven by facies change and taxon rarity and those that represent true regional extinction. This new approach to confidence limits could also be applied to test the synchroneity of extinction or origination at large-scale turnover events, such as mass extinctions and the turnover pulses that bound episodes of faunal stasis.
Shell growth and morphogenesis were studied in nine species of Bivalvia from the viewpoint of theoretical morphology. The aperture map, or pattern of relative rate of shell accretion for each point around the aperture, received particular attention. Morphometric analyses indicate that the basic pattern of the aperture map is generally maintained throughout ontogeny, whereas both shell convexity and aperture shape commonly change with growth. Computer simulations show that posterior elongation of the aperture with growth cancels the effect of ontogenetic shell inflation to move the maximal growth point anteriorly. In the species examined, the coiling axis is generally inclined to the hinge axis toward the anterodorsal direction and is plunging to the dextral side of the valve. This condition allows ontogenetic shell inflation without modification of the basic pattern of the aperture map. The result indicates that ontogenetic change of shell form is architecturally constrained by a basic pattern of the aperture map, which is kept throughout ontogeny.
We examine patterns of intra-otolith variation in δ13C values of fossil Aplodinotus grunniens (freshwater drum) otoliths recovered from an archeological site in northeast Tennessee. We find three repeatable patterns: an initial increase early in ontogeny followed by relatively stable δ13C values as the fish ages, an initial strong covariation between seasonal δ18O and δ13C values, and a decrease with age in the magnitude of seasonal change in δ13C values. These last two observations are illustrated by seasonal least-squares linear regressions between δ13C and δ18O values that tend to progressively decrease in r2 value and slope with fish age. These patterns are evaluated by using a mass balance model in which otolith δ13C values are derived from dissolved inorganic carbon of ambient water mixing with carbon derived from metabolic processes. The proportion of metabolically derived carbon is found to be the dominant factor controlling intra-otolith variation in δ13C values.
Thus, the difference between maximum and minimum δ13C values from a single otolith (δ13Cmax–min) is postulated to reflect the total change in metabolic rate over the lifetime of a fish. δ13Cmax–min values significantly and negatively covary with average δ18O(CaCO3) values, suggesting either a higher total change in metabolic rate over the lifetime of a fish in cooler climates characterized by shorter growing seasons, or a decrease in summer/winter precipitation ratio. A proxy for metabolic rate preserved in otoliths would facilitate the understanding of evolutionary history in physiological traits of fishes and improve our understanding of bioenergetics.
The Desmostylia, an extinct order of mammals related to sirenians and proboscideans, are known from the late Oligocene to late Miocene of the North Pacific. Though often categorized as marine mammals on the basis of fossil occurrences in nearshore deposits, reconstructions of desmostylian habitat and dietary preferences have been somewhat speculative because morphological and sedimentological information is limited. We analyzed the carbon, oxygen, and strontium isotope compositions of enamel from Desmostylus and co-occurring terrestrial and marine taxa from middle Miocene sites in California to address the debate surrounding desmostylian ecology. The δ13C value of tooth enamel can be used as a proxy for diet. Desmostylus had much higher δ13C values than coeval terrestrial or marine mammals, suggesting a unique diet that most likely consisted of aquatic vegetation. Modern aquatic mammals tend to exhibit lower variability in δ18O values than terrestrial mammals. Both fossil marine mammals and Desmostylus exhibited low δ18O variability, suggesting that Desmostylus spent a large amount of time in water. Finally, the Sr isotope composition of marine organisms reflects that of the ocean and is relatively invariant when compared with values for animals from land. Sr isotope values for Desmostylus were similar to those for terrestrial, rather than marine, mammals, suggesting Desmostylus was spending time in estuarine or freshwater environments. Together, isotopic data suggest that Desmostylus was an aquatic herbivore that spent a considerable portion of its life foraging in estuarine and freshwater ecosystems.
The Mesozoic marine revolution focuses on increased predation by durophagous (shell-crushing) predators and the concomitant evolution of prey organisms that occurred in the Mesozoic. Evidence of this predator/prey revolution is found in the appearance and increase of new types of predators that can crush hard shells of prey organisms, and is also found in the morphological changes of prey organisms, such as the appearance of a protective shell morphology of gastropods. We present new data based on the occurrence of shell fragments that indicate a slower increase in durophagous predation than has been considered previously.
The results of an experiment on shell abrasion, in which shells were tumbled in barrels with sediments, indicate that incomplete bivalves and gastropods with angular margins from shallow-marine deposits can be considered as good evidence of durophagous predation. Such angular shell fragments are virtually absent from Japanese Mesozoic shell beds, whereas they are occasionally or commonly found in the Paleogene and are usually abundant in Neogene shell beds. The dominant occurrence of fossil shell fragments in the Cenozoic, as well as the data from shell abrasion experiments using tumbling barrels, indicates that wave agitation or currents do not produce shell fragments with angular margins. Such angular shell fragments are interpreted as the result of durophagous predation that has increased during Cenozoic time, and can be a useful tool in estimating durophagous predation in the fossil record. Revised data on the number of durophagous predator taxa (crustaceans and teleostean fishes) also support this conclusion.
Ediacaran fossils at Mistaken Point, southeastern Newfoundland (terminal Neoproterozoic; 565–575 Ma) represent the oldest known animal communities. In contrast to most Phanerozoic fossil assemblages, in which postmortem transportation, bioturbation, and the accumulation of hardparts obscure community relationships, all fossils in the Mistaken Point assemblages were sessile, soft-bodied organisms that show no evidence of mobility in life or transportation after death. Mistaken Point assemblages are spectacularly preserved on large bedding planes as in situ census populations of hundreds to thousands of fossils, recording the living soft-bodied benthic community at the moment it was smothered by volcanic ash. This unique preservation style allows ecological tests routinely conducted in modern communities (e.g., species richness, abundance, “biomass,” diversity, and evenness, as well as statistical tests of nearest-neighbor interactions) to be applied to the fossil communities. Observed patterns of community variability are consistent with the theory that Mistaken Point fossil surfaces are “snapshots” recording different stages of ecological succession, progressing from communities of low-level feeders (e.g., pectinates and spindles) to frond-dominated communities with complex tiering and spatial structure. The presence of diverse slope communities at Mistaken Point suggests that the deep sea was colonized rapidly during the evolution of complex organisms. Species richness, abundance, and diversity values, as well as levels of intraspecific interaction, all fall within the typical range observed in modern slope communities. These structural similarities imply that ecological processes present in Ediacaran communities at Mistaken Point were strikingly similar to the processes that operate in modern deep-sea animal communities.
Long-term diversity equilibria, ecological incumbency, and widespread recurrent fossil assemblages have each been cited as evidence that local processes, such as competition, played an important role in structuring communities over geologic time. We analyze the relationship between local and regional diversity in tropical marine communities spanning approximately 13 Myr of the Late Ordovician to test for the role of local processes in structuring local communities. We find a significant and strong positive relationship between local and regional diversity, indicating that local communities were not saturated with species and that local processes did not exert a dominant influence on local diversity. Rather, local diversity was influenced more by regional oceanographic processes that governed the size of the regional species pool. This evidence for unsaturated communities is consistent with the Walker and Valentine hierarchically structured niche model of global diversification. These results come at the beginning of the 200-Myr Paleozoic plateau in both local and global diversity and therefore raise the question whether local communities were ever saturated with species during the Paleozoic. Similar studies need to be conducted during other times in the Paleozoic to determine if this is indeed the case.
The asphalt deposits of Rancho La Brea are well known for preserving a prolific and diverse Late Pleistocene fauna. However, little taphonomic research has been done on these collections. To better understand the formation of this impressive assemblage, a taphonomic study of the bones of the large mammals from one asphalt deposit, Pit 91, was carried out, and results are presented here. The predominance of carnivore specimens in the Rancho La Brea deposits has long been explained by a scenario in which a prey animal was trapped in asphalt and attracted large numbers of carnivores who became trapped in turn. Hypotheses generated from this scenario were tested by collecting taphonomic data on over 18,000 specimens. Weathering data indicate that elements were deposited fairly rapidly. However, patterns of skeletal part representation for the seven most common species demonstrate that complete skeletons are not present. Water transport is ruled out as the primary process responsible for removing skeletal elements based on abrasion data. Instead, the feeding activity of carnivores (ravaging) appears to have been an important factor in the formation of the assemblage.
Bivalves and gastropods, prominent members of the Modern Evolutionary Fauna, are traditionally noted for sharing remarkably similar global diversity trajectories and environmental distributions throughout the Phanerozoic. By comparing their fossil occurrences at several scales within a finely resolved geographic, environmental, and temporal framework, it is possible to evaluate whether such similarities are caused primarily by intrinsic macroevolutionary factors or extrinsic ecological factors. Using a database of 7779 global gastropod and bivalve genus occurrences, we investigate the geographical and environmental attributes of bivalves and gastropods during the Ordovician Period at scales ranging from global, to a comparison among five paleocontinents, to an intracontinental comparison of four regions within Laurentia. Although both classes shared statistically indistinguishable global diversity trajectories and broadly similar environmental distributions during the Ordovician, their environmental distributions differed in several significant features. Furthermore, the diversity trajectories and environmental distributions of these classes differed significantly among paleocontinents and among regions within Laurentia. Bivalves were consistently most diverse in deeper water, siliciclastic-rich settings in higher-latitude paleocontinents whereas gastropods were consistently most diverse in shallower, carbonate-rich settings in more-equatorial paleocontinents. Notably, these environmental differences were robust to changing physical parameters within paleocontinents, with each class consistently tracking its preferred environmental setting. These results suggest that environmental factors played significant, albeit distinct, roles in the Ordovician diversifications of gastropods and bivalves. However, their similar global diversity trajectories suggest that shared, intrinsic macroevolutionary attributes also may have played an important role in the evolution of these classes during the Ordovician Radiation.
The single bony element forming the lower jaw of living mammals, the dentary, has been interpreted as representing the culmination of a long and gradual evolutionary trend. Numerous fossil nonmammalian synapsids (“mammal-like reptiles”) show varying degrees of enlargement of the dentary and concomitant reduction in the postdentary bones. To quantitatively reexamine patterns of morphological change in the evolution of the mammalian lower jaw, measurement and discrete character data were collected from 322 fossil synapsid mandibles spanning Late Carboniferous through Jurassic time. Measurements confirm that the relative contribution of the dentary increased in theriodont (advanced therapsid) evolution with regard to both stratigraphic and phylogenetic position. However, dentary enlargement and postdentary reduction failed to typify all therapsid subclades. Qualitative characters of the mandible were used to quantify morphological similarity with regard to the early mammal Morganucodon. Analyses contrasting stratigraphic and phylogenetic position with mammalian similarity indicate that mandibular evolution was primarily conservative, with only anomodont therapsids evolving substantial morphological novelty. Scaling analyses comparing the area of the dentary and postdentary regions to jaw length uniformly show isometry or slight positive allometry, although cynodont therapsids have a smaller postdentary region than any other therapsid subgroup. These results suggest that body size decreases cannot fully explain the reduction of the postdentary bones. Finally, step size bias was tested as a mechanism for explaining long-term trends. Qualitative data reveal no significant difference in the magnitude of character changes occurring in mammalian and nonmammalian directions.